Hemerotrecha banksi (Arachnida, Solifugae), a diurnal group of solifuges from North America

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Hemerotrecha banksi (Arachnida, Solifugae), a diurnal group of solifuges from North America Jack O. Brookhart and Paula E. Cushing: Department of Zoology, Denver Museum of Nature and Science, 2001 Colorado Boulevard, Denver, Colorado 80205-5798, USA. E-mail: [email protected] Abstract. The Hemerotrecha banksi group is revised and the status of the genus Hemerotrecha is examined. The female of H. truncata Muma 1951 is described for the first time, and five new species are named: H. hanfordana, H. kaboomi, H. prenticei. H. pseudotruncata, and H. vetteri. Keywords: Solifuges, new species, distribution, systematics Although solifuges are considered mainly nocturnal (Muma 1951, 1970; Cloudsley-Thompson 1958; Lawrence 1960, 1962; Punzo 1998), many species are primarily diurnal including species in the South African Solpugidae (Wharton 1981, 1987) and South American members of the Mummuciidae (Maury 1985; Xavier & Rocha 2001). The North American banksi group of the Eremobatidae genus Hemerotrecha also appears to be diurnal. The Greek Hemerous, the sun, and trechos, to run, were used by Banks (1903) to erect the genus Hemerotrecha. He designated Hemerotrecha californica Banks 1903 as the type species. The collector, Dr. Harold Heath, said, ‘‘they run about in the blazing hot sunshine’’ (Banks 1903). The genus Hemerotrecha currently includes 31 species from western United States and northern Mexico (Harvey 2003). As defined by Muma (1951, 1970), the genus Hemerotrecha includes small to moderate-sized solifuges bearing a style-like fixed cheliceral finger with the ventral edge irregularly undulate or bearing one or more modified teeth. The mesoventral groove is absent or, at most, very faint. The flagellum complex consists of a dorsal row of striate bristles, the striae formed by very tiny setae, and a ventral row of plumose setae that are more plumose apically (Fig. 1). Muma (1951) established the Hemerotrecha banksi group for four species all with characters so similar that he thought that they might in fact be one species. The group includes: H. banksi Muma 1951, the new name for H. californica Banks 1903 (which is a junior secondary homonym of Cleobis californica Banks 1899); H. californica (Banks 1899); H. marginata (Kraepelin 1911); and H. truncata Muma 1951. All four species have eyes separated by 1.5–2 diameters, males with the ventral margin of the fixed cheliceral finger irregularly undulate, striate bristles of the flagellum complex indistinctly striate with apical and sub- apical bristles broad and flattened, and females with roughly triangular genital opercula, with parallel medial margins. Muma’s 1970 publication emphasized the shape of the tip of the male fixed finger, length and type of ctenidia, and palpal coloration. In addition we found them to possess dark edges on the anterior of the malleoli in both males and females (Fig. 2), and a cup-like mesoventral groove at the tip of the male fixed finger (arrow, Fig. 3). Females were described for H. banksi and H. californica but not for H. truncata. Muma did not examine Kraepelin’s types of Eremobates marginatus Kraepelin 1911 but used Roewer’s (1934) drawings. The type localities of each species were in the state of California and the meager material examined was all from that state except for a male and female from the state of Washington. Muma (1962, 1963, 1970) included specimens from these states as well as Idaho, Oregon, and several specimens that he identified as Hemerotrecha californica from the Nevada Test Site. After examining more extensive solifuge collections, particularly those of D. Giuliani (CAS, ESS) including specimens from wide ranging regions of Southern California and Arizona, a collection by Wendel Icenogle of Winchester, California, a collection from Tom Prentice (UCR) and a group sent to us by Richard Zack collected at the Hanford Nuclear Reactor Site, Hanford, Washington, we decided that a revision of the group was in order. METHODS Using the methods of Muma (1951), Brookhart & Muma (1981, 1987), Muma & Brookhart (1988), and Brookhart & Cushing (2004), we measured total body length, length of palpus, leg I, and leg IV of both males and females. Length and width of chelicera and propeltidium, width of base of fixed finger, and genital operculum were all measured using Spot Basic TM on an Olympus SZX12. All measurements are in millimeters. Ratios used previously by Muma (1951, 1970, 1989), Brookhart & Muma (1981, 1987), Muma & Brookhart (1988), and Brookhart & Cushing (2002, 2004) were comput- ed. These ratios are as follows: A/CP: the sum of the lengths of palpus, leg I, and leg IV divided by the sum of length of chelicera and propeltidium indicating length of appendages in relation to body size. Long-legged species have larger A/CP ratios. Because there is no fondal notch, the chelicera width/ fixed finger width ratio is used to indicate whether the fixed cheliceral finger is thin or robust in relation to the size of the chelicera. Genital operculum length/genital operculum width demonstrates the relative size of the female genital operculum in terms of length and width. No statistical analysis was attempted because of small sample sizes. Species determinations were based on a combination of color comparisons, general shape of male fixed finger, particularly the tip; palpal setation; ctenidial size and shape. The difference (or similarity) in color pattern between the chelicerae and the propeltidium was noted as was the color pattern of the palpus and legs. The shape of the female chelicera and the female genital operculum margin were observed using the method of Brookhart & Cushing (2004). Due to strong similarities among members of this group, the description of H. banksi was used as the basis for all other species descriptions. Female morphology is consistent among the various species; differences are described when necessary. 2008. The Journal of Arachnology 36:49–64 49

Transcript of Hemerotrecha banksi (Arachnida, Solifugae), a diurnal group of solifuges from North America

Hemerotrecha banksi (Arachnida, Solifugae), a diurnal group of solifuges from North America

Jack O. Brookhart and Paula E. Cushing: Department of Zoology, Denver Museum of Nature and Science, 2001

Colorado Boulevard, Denver, Colorado 80205-5798, USA. E-mail: [email protected]

Abstract. The Hemerotrecha banksi group is revised and the status of the genus Hemerotrecha is examined. The female ofH. truncata Muma 1951 is described for the first time, and five new species are named: H. hanfordana, H. kaboomi, H.prenticei. H. pseudotruncata, and H. vetteri.

Keywords: Solifuges, new species, distribution, systematics

Although solifuges are considered mainly nocturnal (Muma1951, 1970; Cloudsley-Thompson 1958; Lawrence 1960, 1962;Punzo 1998), many species are primarily diurnal includingspecies in the South African Solpugidae (Wharton 1981, 1987)and South American members of the Mummuciidae (Maury1985; Xavier & Rocha 2001). The North American banksigroup of the Eremobatidae genus Hemerotrecha also appearsto be diurnal.

The Greek Hemerous, the sun, and trechos, to run, were usedby Banks (1903) to erect the genus Hemerotrecha. He designatedHemerotrecha californica Banks 1903 as the type species. Thecollector, Dr. Harold Heath, said, ‘‘they run about in theblazing hot sunshine’’ (Banks 1903). The genus Hemerotrechacurrently includes 31 species from western United States andnorthern Mexico (Harvey 2003). As defined by Muma (1951,1970), the genus Hemerotrecha includes small to moderate-sizedsolifuges bearing a style-like fixed cheliceral finger with theventral edge irregularly undulate or bearing one or moremodified teeth. The mesoventral groove is absent or, at most,very faint. The flagellum complex consists of a dorsal row ofstriate bristles, the striae formed by very tiny setae, and a ventralrow of plumose setae that are more plumose apically (Fig. 1).

Muma (1951) established the Hemerotrecha banksi group forfour species all with characters so similar that he thought thatthey might in fact be one species. The group includes: H. banksiMuma 1951, the new name for H. californica Banks 1903 (whichis a junior secondary homonym of Cleobis californica Banks1899); H. californica (Banks 1899); H. marginata (Kraepelin1911); and H. truncata Muma 1951. All four species have eyesseparated by 1.5–2 diameters, males with the ventral margin ofthe fixed cheliceral finger irregularly undulate, striate bristles ofthe flagellum complex indistinctly striate with apical and sub-apical bristles broad and flattened, and females with roughlytriangular genital opercula, with parallel medial margins.Muma’s 1970 publication emphasized the shape of the tip ofthe male fixed finger, length and type of ctenidia, and palpalcoloration. In addition we found them to possess dark edges onthe anterior of the malleoli in both males and females (Fig. 2),and a cup-like mesoventral groove at the tip of the male fixedfinger (arrow, Fig. 3).

Females were described for H. banksi and H. californica butnot for H. truncata. Muma did not examine Kraepelin’s typesof Eremobates marginatus Kraepelin 1911 but used Roewer’s(1934) drawings. The type localities of each species were in thestate of California and the meager material examined was allfrom that state except for a male and female from the state of

Washington. Muma (1962, 1963, 1970) included specimensfrom these states as well as Idaho, Oregon, and severalspecimens that he identified as Hemerotrecha californica fromthe Nevada Test Site. After examining more extensive solifugecollections, particularly those of D. Giuliani (CAS, ESS)including specimens from wide ranging regions of SouthernCalifornia and Arizona, a collection by Wendel Icenogle ofWinchester, California, a collection from Tom Prentice (UCR)and a group sent to us by Richard Zack collected at theHanford Nuclear Reactor Site, Hanford, Washington, wedecided that a revision of the group was in order.

METHODS

Using the methods of Muma (1951), Brookhart & Muma(1981, 1987), Muma & Brookhart (1988), and Brookhart &Cushing (2004), we measured total body length, length ofpalpus, leg I, and leg IV of both males and females. Lengthand width of chelicera and propeltidium, width of base offixed finger, and genital operculum were all measured usingSpot BasicTM on an Olympus SZX12. All measurements are inmillimeters. Ratios used previously by Muma (1951, 1970,1989), Brookhart & Muma (1981, 1987), Muma & Brookhart(1988), and Brookhart & Cushing (2002, 2004) were comput-ed. These ratios are as follows: A/CP: the sum of the lengths ofpalpus, leg I, and leg IV divided by the sum of length ofchelicera and propeltidium indicating length of appendages inrelation to body size. Long-legged species have larger A/CPratios. Because there is no fondal notch, the chelicera width/fixed finger width ratio is used to indicate whether the fixedcheliceral finger is thin or robust in relation to the size of thechelicera. Genital operculum length/genital operculum widthdemonstrates the relative size of the female genital operculumin terms of length and width. No statistical analysis wasattempted because of small sample sizes.

Species determinations were based on a combination ofcolor comparisons, general shape of male fixed finger,particularly the tip; palpal setation; ctenidial size and shape.The difference (or similarity) in color pattern between thechelicerae and the propeltidium was noted as was the colorpattern of the palpus and legs. The shape of the femalechelicera and the female genital operculum margin wereobserved using the method of Brookhart & Cushing (2004).Due to strong similarities among members of this group, thedescription of H. banksi was used as the basis for all otherspecies descriptions. Female morphology is consistent amongthe various species; differences are described when necessary.

2008. The Journal of Arachnology 36:49–64

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Specimens examined for this study are deposited inthe American Museum of Natural History, New York(AMNH); Brigham Young University, Provo, Utah (BYU);California Academy of Sciences, San Francisco (CAS);Denver Museum of Nature & Science, Denver, Colorado(DMNS); Essig Museum, University of California at Berkeley,Berkeley (ESS); University of California at Riverside (UCR);California State University, Northridge (CSN); FloridaState Collections of Arthropods, Gainesville (FSCA); Muse-um of Comparative Zoology, Cambridge, Massachusetts(MCZ); San Diego Museum of Natural History, San Diego,California (SMNH); Washington State University, Pullman(WSU); and Zoologische Staatssammlung, Munich, Germany(ZSM).

SYSTEMATICS

Family Eremobatidae Kraepelin 1901

Subfamily Therobatinae Muma 1951

Genus Hemerotrecha Banks 1903

Type species.—Hemerotrecha californica Banks 1903 (juniorsecondary homonym of Cleobis californica Banks 1899, nowHemerotrecha banksi Muma 1951), by monotypy.

Remarks.—Muma (1951) placed the genus Hemerotrechain the Therobatinae based on the style-like fixed cheliceralfinger, undulate ventrally or with small modified denticles,none, or, at best, a very faint mesal ventral groove, andthe female operculum variously developed. He latermodified his description of the genus Hemerotrecha (Muma1970) by describing the fixed cheliceral finger as weaklycurved, undulate, or turned downward at the tip, and thedorsal flagellar setae as striate, spatulate, or hooked.

Muma (1989) refined his description of the fixed fingeras essentially straight but at times being denticulate,undulate, or serrate and completely lacking a mesal ormesoventral groove, the flagellar setae additionally to include‘‘strong, enlarged, flattened, or hooked’’ setae, and thefemale opercula being variable but consistent within speciesgroups.

KEY TO THE MALES OF THE HEMEROTRECHABANKSI GROUP

1 Chelicerae and propeltidium of the same coloration . . 2

Chelicerae paler than propeltidium . . . . . . . . . . . . . . 5

2 Chelicera and propeltidium pale yellow; eye tubercle pale;

cheliceral tip of fixed finger rounded (Fig. 28); palpus

dusky on metatarsus and distal 20% of tarsus (Fig. 27);

ctenidia short, thin, pointed . . . . Hemerotrecha marginata

Chelicera and propeltidium dark to dusky; eye tubercle

dark; cheliceral tip hooked (Fig. 45); palpal tarsus either

pale or completely dark; ctenidia thin or broad extending

from one half to the entire length of the succeeding

sternite (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3 Entire body dark; entire palpus (Fig. 46) and appendages

dark; chelicera slightly hooked with a distinct ridge on the

distal dorsal edge of the fixed finger (arrow, Fig. 44)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemerotrecha vetteri

Chelicera and propeltidium blotchy gray or blotchy

orange (Fig. 15); palpal metatarsus dark (Fig. 18) . . . . 4

4 Cheliceral fixed finger slightly hooked with tiny ridge on the

distal dorsal edge of fixed finger, a short, deep concavity on

the apical ventral edge of fixed finger (Fig. 16), ctenidia thin

and of medium length . . . . . . . . . . Hemerotrecha hanfordana

Cheliceral fixed finger slightly hooked without a distal

dorsal ridge or an apical, ventral concavity (Fig. 51),

ctenidia short and broad (Fig. 52) . . Hemerotrecha prenticei

5 Fixed cheliceral finger gently curved dorsally with

rounded, attenuated tip (Fig. 12) . . . . . . . . . . . . . . . . 6

Fixed cheliceral finger gently curved dorsally with slightly

to strongly hooked tip (Fig. 38) . . . . . . . . . . . . . . . . . 7

6 Palpus dusky on metatarsus (Fig. 11); 2 long ctenidia

extending length of sternal segment (Fig. 13); body pale;

without spine-like setae on ventral tarsus and metatarsus

of palp . . . . . . . . . . . . . . . . . . . Hemerotrecha californica

Palpus dusky on tarsus and metatarsus (Fig. 7); 2 shorter

ctenidia (Fig. 6); body dark; with spine like setae on ventral

tarsus and metatarsus of palp (Fig. 7) . . Hemerotrecha banksi

7 Fixed cheliceral finger with apical hook but with

otherwise typical curved shape (Fig. 22); dusky orange

chelicera; blotchy, dark propeltidium (Fig. 21); creamy

yellow abdomen . . . . . . . . . . . . . . . Hemerotrecha kaboomi

Fixed cheliceral finger attenuated with a noticeable hook

(Figs. 32, 38); dark to dusky orange chelicera and

propeltidium, dusky to dark abdomen . . . . . . . . . . . . 8

8 Fixed cheliceral finger with sharply hooked (parrot beak)

tip (Fig. 32); dusky chelicera, blotchy dark propeltidium

(Fig. 31) and abdomen; dusky palpal tarsus, metatarsus,

tibia and tip of femur . . . . . . . . . . . Hemerotrecha truncata

Fixed cheliceral finger with hooked tip but not nearly as

deep or sharp as above (Fig. 38); dusky chelicera; blotchy

propeltidium (Fig. 37); creamy yellow abdomen; dusky

palpal metatarsus (Fig. 39) . . Hemerotrecha pseudotruncata

Figures 1–3.—Hemerotrecha vetteri and H. banksi. 1. Ectal view ofH. hanfordana chelicera showing flagellum complex. 2. Ventral viewof malleoli of Hemerotrecha vetteri, new species showing dark edges.3. Ectal view of the right chelicera of Hemerotrecha banksi holotypeshowing mesal ventral groove (arrow). Scale bars 5 1 mm.

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Hemerotrecha banksi Muma 1951Figs. 3–9, 54

Hemerotrecha californica Banks 1903:79 (junior secondaryhomonym of Cleobis californica Banks 1899).

Hemerotrecha banksi Muma 1951:99–100, figs. 185–192(replacement name for Hemerotrecha californica Banks1903).

Material examined.—Types: USA: California: male holo-type, Monterey County, Pacific Grove (36.62uN, 121.92uW),no date, H. Heath (MCZ). ‘‘Allotype’’ (designated by Muma1951), 1 female, San Mateo County, Redwood City (37.49uN,122.24uW), 18 May 1924, W. Meehan (AMNH).

Other material examined: USA: California: CalaverasCounty: 1 L, West Point (38.4uN, 120.53uW), 7 August 1970,S.C. Williams (CAS); Contra Costa County: 2 L, 1 K,Redwood (37.49uN, 122.29uW), 14 April 1995, J.G. Rozen,J.W. McSwain (ESS); 1 L, Walnut Creek (37.90uN, 122.06uW),3 June 1961, J. Powell (CAS); Marin County: 1 L, Tiburon,Ring Mountain (37.92uN, 122.49uW), 13 July 1977, T.S.Briggs (ESS); Monterey County: 1 L, Pacific Grove (36.62uN,121.92uW), no date, E.C. Starks (AMNH); 1 K, Redwood City(37.49uN, 122.24uW), 18 May 1962, W. Meehan (AMNH);San Benito County: 1 L, 9.7 km SE. of Idria (36.42uN,120.63uW), 29 June 1954, S.G. Rozen (CAS); 1 L, PinnacleNational Monument (36.47uN, 121.17uW), 7 May 1977 C.E.Griswold (CAS); San Mateo County: 1 L, 19 April 1918, B.H.Van Duzen (CAS); Sonoma County: 1 L, Petaluma (38.26uN,122.53uW), 4 July 1979, D.H. Kavanaugh (CAS); Santa ClaraCounty: 1 K, Jasper Ridge (37.41uN, 122.27uW), 31 October1952, F.S. Bartholomew (CAS); Santa Cruz County: 1 L,Scotts Valley (37.05uN, 122.01uW), 22 May 1990, running ontrail, R. Morgan (CAS); 1 L, Santa Cruz Grasslands(coordinates not determined), 22 May 1990, R. Morgan(CAS); Santa Margarita County: 1 L, 9.7 km E. of San LuisObispo (35.28uN, 120.66uW), 11 June 1958, J. W. McSwain(ESS).

Diagnosis.—Tip of male fixed cheliceral finger rounded,tapering anteriorally, palpal tarsus dark for at least the distalhalf, metatarsus dark 80% apically, dusky creamy orangechelicera, dark to dusky light orange propeltidium; ctenidiashort (about half the succeeding segment), pointed. Femaleswith same coloration.

Description.—Male, Color: chelicera dusky yellow-orange,propeltidium dusky to dark orange but always darker thanchelicera (Fig. 4), eye tubercle dark, abdomen blotchy, duskyto dark, palpal tarsus and apical 80% metatarsus dusky todark; leg I dark on tarsi and metatarsi, legs II, III, IV darkventrally.

Chelicera: fixed finger with a small basal rise and a roundedtip, ventral edge straight (although two specimens had a slightventral concavity) with no apparent hooklike terminus; themesal ventral groove a small, shallow apical cup; movablefinger with large primary tooth, smaller anterior tooth, 1–2intermediate teeth, the posterior being separated from primarytooth. Small crenulations anterior to anterior tooth. No mesaltooth (Fig. 5). Fondal notch obscure to absent, fondal teethgraded I, III, II, IV. Apical, subapical plumose flagellarbristles as well as dorsal apical cheliceral setae flattened(Fig. 5, arrow).

Propeltidium: eyes separated by 1.5 diameter of eye (Fig. 4).

Abdomen: 2 short, pointed ctenidia ventrally on fourthabdominal sternite (Fig. 6). Edges of malleoli darkly tinged.

Palp: scopula absent; a row of 2–3 spine-like setae on mesalventral portion of metatarsus (Fig. 7).

Dimensions: male holotype: total length 11.0, chelicerallength 2.6, cheliceral width 1.2, propeltidium length 1.5,propeltidium width 2.0, palpus length 7.2, first leg length 5.0,fourth leg length 12.1. Ratios: A/CP 5.92, cheliceral width/fixed finger width 4.28.

Male (5): total length 8.5–11.0, cheliceral length 2.13–2.96,cheliceral width 0.93–1.28, propeltidium length 1.09–1.45,propeltidium width 1.71–2.23, palpus length 6.5–7.8, first leglength 4.6–7.6, fourth leg length 9.0–13.5. Ratios: A/CP 5.07–6.99, cheliceral width/fixed finger width 3.55–4.23.

Female: Color: same as males.

Chelicera: fixed finger with large primary tooth and medialtooth, smaller anterior tooth. Two intermediate teeth betweenprimary tooth and medial tooth and medial tooth and anteriortooth. movable finger with medium sized primary tooth andanterior tooth. Two intermediate teeth between primary toothand anterior tooth, the posterior intermediate tooth in thenotch of the primary tooth. No mesal tooth (Fig. 8).

Genital region: genital operculum roughly triangular withparallel median margins, no wings, and a gently curvedposterior margin. Genital opening behind the opercular plates(Fig. 9).

Dimensions: Female allotype: total length 11.0, chelicerallength 4.1, cheliceral width 1.6, propeltidium length 1.6,propeltidium width 2.7, palpus length 6.5, first leg length 5.5,fourth leg length 9.0. Ratios: A/CP 3.7, genital operculumlength/genital operculum width 1.5.

Female measurements (n 5 3): total length 9.0–12.0,cheliceral length 2.23–2.28, cheliceral width 1.12–1.22, pro-peltidium length 1.38–1.4, propeltidium width 1.82–2.0, palpuslength 5.5–8.0, first leg length 4.3–5.5, fourth leg length 8.9–9.7. Ratios: A/CP 5.06–5.27, genital operculum length/genitaloperculum width 1.5–1.52.

Remarks.—The distribution of this species roughly encom-passes the northern area of California surrounding andincluding the Monterey Peninsula (Fig. 54). Muma (1951)lists female paratypes deposited at Cornell University andUtah State University (USU) but USU material is now atAMNH and the Cornell material is missing.

Hemerotrecha californica (Banks 1899)

Figs. 10–14, 54

Cleobis californica Banks 1899:314–315.

Ammotrecha californica (Banks): Banks 1900:427.

Hemerotrecha californica (Banks): Banks 1904:363.

Material examined.—Type: USA: California: female holo-type, Los Angeles, Los Angeles County, A. Davidson (MCZ).

Other material: USA: Arizona: Mohave County: 7 L, 2 K,Virgin River Canyon, 11.3 km E. of Littlefield (36.52uN,113.55uW), March-September 1983, D. Giuliani (ESS).California: Inyo County: 1 L, 30.6 km N. of Ridgecrest(35.62uN, 117.97uW), 5 April 1983–13 September 1983, D.Giuliani (ESS); 1 L, Saline Valley, Granite Canyon (35.82 uN,116.62uW) 25 May 1981–15 April 1982, D. Giuliani (ESS); 2 L,White Mountains, Big Pine (37.09uN, 118.19uW), 1 November1985–28 June 1986, D. Giuliani, (ESS); 8 L, 2 K, Sierra Nevada

BROOKHART & CUSHING—HEMEROTRECHA BANKSI GROUP (SOLIFUGAE) 51

Range, 12.9 km NE. of Big Pine (37.09uN, 118.19uW), May–November 1983, D. Giuliani (ESS); 2 L, Bishop (37.21uN,118.25uW), October 1985–November 1986, D. Giuliani (ESS);Los Angeles County: 1 L, Campo (34.14uN, 118.36uW), 23October 1970, C. Mahrdt (CAS); 1 L, same data except 11 July1971 (CAS); 1 L, same data except 31 July 1971 (CAS); 1 K,same data except 31 July 1971, H. Sunclar (CSN); 1 L, SanPedro (33.85uN, 118.31uW), (no date), Banks type label (MCZ#14855); 1 L, Chatsworth (34.26uN, 118.60uW), 1 July 1962,W.E. Icenogle (CSN); San Bernardino County: 2 L CovingtonFlats (34.04uN, 116.31uW), June 2001, USGS, San Diego(DMNS), 1 L, Winchester (33.37uN, 17.15uW), 15 June 1967,W.R. Icenogle (DMNS); 3 L, same data except 29 June 1967(DMNS); 1 K, same data except 5 May 1968 (DMNS); 1 L,same data except 6 June 1968 (DMNS); 1 L, same data except

21 May 1969 (DMNS); 1 L, same data except 10 May 1970(DMNS). Nevada: Clark County: 2 L, 1 K, 14.5 km SW. ofOverton (36.51uN, 114.32uW), March-September 1983, D.Giuliani (ESS); 3 L, 1 K, Spring Range Canyon (36.152uN,115.88uW), March-September 1983, D. Giuliani (ESS). NewMexico: San Juan County: 4 L, 1 K, Shiprock (36.79uN,108.69uW), March-September 1984, D. Giuliani (ESS).

Diagnosis.—Chelicera and propeltidium both dusky yellow-orange. Palpus dark on apical 80% of metatarsus. Fixedcheliceral finger slightly curved dorsally with a cuppedmesoventral groove apically. Two long ctenidia extendinglength of succeeding sternite, palpus with no spine-like setaeon ventral side.

Description.—Male (from Winchester, California), Color:chelicera and propeltidium dusky yellow to dusky orange,

Figures 4–9.—Hemerotrecha banksi Muma. 4–7. Male holotype: 4. Dorsal view, propeltidium; 5. Mesal view, right chelicera showing flagellacomplex (arrow); 6. Ventral view, ctenidia; 7. Mesal ventral view, right palpus. 8–9. Female allotype: 8. Ectal view, chelicera; 9. Ventral view,genital operculum. Scale bar 5 1 mm.

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propeltidium dusky orange to tan (Fig. 10), eye tubercle dark;abdomen lighter centrally with dusky tan blotches ectally,palpal metatarsus dark to dusky on apical 80% (Fig. 11); leg Ipale, legs II, III, IV faintly tan dorsally.

Chelicera: fixed finger with a shallow cup-like mesal ventralgroove apically but otherwise normally rounded at the tip; nodorsal ridge, movable finger with a more flattened, triangularshaped anterior tooth; 2 intermediate teeth, posterior inter-mediate tooth separated from primary tooth; a small butdiscernable fondal notch (Fig. 12); flagellar complex typical ofthe group.

Abdomen: two ctenidia extending length of succeedingsternite (Fig. 13). Edges of malleoli darkly tinged.

Palp: many hair-like setae but no spines (Fig. 11).Dimensions: (n54): total length 8.0–9.0, cheliceral length

2.22–4.0, cheliceral width 0.95–1.9, propeltidium length 1.22–1.3, propeltidium width 1.76–2.1, palpus length 6.0–8.0, firstleg length 4.8–7.0, fourth leg length 8.5–10.5. Ratios: A/CP4.8–6.1, cheliceral width/fixed finger width 3.9–4.4.

Females.—Color: holotype lighter than male description butmost other females the same coloration as males.

Chelicera: similar to H. banksi but posterior intermediatetooth is imperceptively to visibly separated from primarytooth (Fig. 14).

Genital region: typical, perhaps more flattened on theposterior margin.

Female allotype: total length 12, cheliceral length 3.8,cheliceral width 1.4, propeltidium length 1.6, propeltidiumwidth 2.6, palpus length 6.5, first leg length 5.5, fourth leglength 9.5. Ratios: A/CP 4.4, 2.7, 0.62, genital operculumlength/genital operculum width 1.6.

Female measurements (n54): total length 8.0–14.0, chelic-eral length 2.5–3.5, cheliceral width 0.92–1.4, propeltidiumlength 1.31, propeltidium width 1.6–1.91, palpus length 5.2–6.0, first leg length 4.5–5.5, fourth leg length 8.5–12.0. Ratios:A/CP 4.7–4.84, genital operculum length/genital operculumwidth 1.4–1.6.

Remarks.—Muma (1951) provides a description of a maleH. californica. Based on the locality information he providesfor examined specimens, this description was based upon amale collected from Starbucks, Washington. The color patternof the propeltidium and chelicerae indicate that this maleactually represents H. hanfordana, new species. Our identifi-cation for this group is based on the length and shape of themale ctenidia as well as the shape of the male fixed finger,particularly the tip. The distribution of H. californica includesthe coastal region from Los Angeles to San Diego acrosssouthern California, into parts of Nevada and into north-western New Mexico. This is in stark contrast to the patchydistribution of other members of this group. However, thiswide distribution is not unusual among other solifuges, i.e.,Eremobates nodularis Muma 1951, Eremochelis bilobatus

Figures 10–14.—Hemerotrecha californica (Banks). 10–13. Male from Winchester, California: 10. Dorsal view, propeltidium; 11. Mesalventral view, right palpus; 12. Ectal view, right chelicera; 13. Ventral view, ctenidia. 14. Female holotype, ectal view, right chelicera. Scale bar 5

1 mm.

BROOKHART & CUSHING—HEMEROTRECHA BANKSI GROUP (SOLIFUGAE) 53

(Muma 1951), Hemerotrecha fruitana Muma 1951 (Brookhart& Brookhart 2006).

Hemerotrecha hanfordana new species

Figs. 15–20, 54

Material examined.—USA: Washington: male holotype,Hanford National Monument, Wahluke Wildlife Area, WhiteBluffs Ferry, Franklin County (46.41uN, 119.47uW) 5–12 July2002, R.S. Zack (DMNS). Female allotype, 2 male and 2female paratypes, collected with holotype (DMNS); 3 male, 1female paratypes (WSU).

Other material: USA: California: Siskiyou County: 2 L,Macdoel (41.83uN, 122.35uW), 6 July 1968, J. Schuh(AMNH); Nevada: Washoe County: 1 L, Wadsworth(39.66uN, 119.29uW), 23 July 1965, B. Opler (ESS); Utah:Box Elder County: 1 L, Lucin (41.35uN, 113.90uW), 19 June1952, D.E. Beck (BYU); 10 L, 5 K, Key Springs Rd. (41.601uN,113.891uW, 2 June–2 July 2007, A. Spriggs & Joey Slowik(DMNS); Tooele County: 1 L, Dugway Proving Grounds(dunes) (44.48uN, 123.36uW), no name (BYU); Washington:Benton County: 8 L, 4 K, Hanford Nuclear Site (46.32uN,119.31uW), Rattlesnake Mountain (46.40uN, 119.61uW), 21June–13 Aug 2002, R.S. Zack (WSU); 1 L, Rattlesnake Spring(46.50uN, 119.71uW), 1–9 July 2002, R.S. Zack (WSU); GrantCounty: 6 L, 4 K, Saddle Mountain National Wildlife Refuge(46.68uN, 119.63uW), 14 June–23 July 2002, R.S. Zack (WSU).

Etymology.—The species name is an adjective referring tothe type locality.

Diagnosis.—Hemerotrecha hanfordana has both darklycolored chelicera and propeltidium; palpus dark on metatar-sus; two short, flat, pointed ctenidia separate it from bothHemerotrecha kaboomi, new species and Hemerotrecha cali-fornica, which have longer, thinner ctenidia. Hemerotrechabanksi also has darkly colored palpal tarsi.

Description.—Male, Color: base coloration a blotchy black,with lighter colored patches on the posterior medianpropeltidium, mesopeltidium, and metapeltidium. Cheliceraand propeltidium both blotchy black (Figs. 15, 16); palpalmetatarsus blotchy black (Fig. 18), leg I dusky on tibia andmetatarsus, legs II, III, IV dark on the dorsal, ectal regions ofcoxa, femur, tibia, tarsus, and metatarsus.

Chelicera: fixed finger with small ridge basally on the dorsaledge, and a slightly hooked tip, ventral edge smooth with ashort, deep mesal ventral groove apically (arrow, Fig. 16).Two specimens had very tiny denticles. Movable finger withlarge primary tooth, two intermediate teeth, and anteriortooth, posterior intermediate tooth is separate from primarytooth and fondal notch obscure to absent; fondal teeth gradedI, III, II, IV (Figs. 16, 17).

Abdomen: two flat, pointed ctenidia extending across halfof the succeeding abdominal segment (Fig. 19).

Palp: palpi with 2–4 thick spine-like setae ventrally(Fig. 18).

Dimensions: Male holotype: total length 11.5, chelicerallength 2.5, cheliceral width 1.1, propeltidium length 1.3,propeltidium width 2.0, palpus length 7.0, first leg length 6.0,fourth leg length 12.5. Ratios: A/CP 6.65, cheliceral width/fixed finger width 4.13.

Male paratypes (5): total length 11.5–13.0, cheliceral length2.33–2.67, cheliceral width 1.05–1.1, propeltidium length 1.33–

1.55, propeltidium width 1.77–2.0, palpus length 6.0–8.0, firstleg length 5.0–6.5, fourth leg length 10.0–12.5. Ratios: A/CP5.59–6.65, cheliceral width/fixed finger width 3.20–4.13.

Females: coloration as in males.

Chelicera: fixed finger with primary tooth, medial tooth,anterior tooth, with two intermediate teeth between each.Movable finger with primary tooth, anterior tooth, and twointermediate teeth. Posterior intermediate tooth in notch ofprimary tooth (Fig. 20).

Genital region: typical configuration for the group.

Dimensions: Female allotype: total length 11.0, chelicerallength 2.67, cheliceral width 0.85, propeltidium length 1.4,propeltidium width 1.85, palpus length 6.0, first leg length 5.0,fourth leg length 8.0. Ratios: A/CP 4.67, genital operculumlength/genital operculum width 2.40. Female paratypes (3):total length 11.0–13.0, cheliceral length 2.7–3.6, cheliceralwidth 0.85–1.46, propeltidium length 1.4–1.87, propeltidiumwidth 1.85–2.6, palpus length 6.0–8.0, first leg length 5.0–7.0,fourth leg length 8.0–14.0. Ratios: A/CP 2.3–4.7, genitaloperculum length/genital operculum width 1.7–2.4.

Remarks.—The male of H. californica described by Muma(1951) was probably a member of this species. Hemerotrechahanfordana is found in xeric regions of the Basin and Rangesystem and extends into both northeastern California andnorthwestern Utah. This roughly corresponds with thedistribution of Eremochelis bidipressus Muma 1951 andanother Hemerotrecha, H. denticulata Muma 1951 (Brookhart& Brookhart 2006).

Hemerotrecha kaboomi new species

Figs. 21–26, 54

Material examined.—Types: USA: Nevada: Nye County:male holotype, Mercury, 25 July 1960, Atomic EnergyCommission and Brigham Young University (AMNH).Female allotype, same data as holotype except 22 June 1960(AMNH). Paratypes: 6 males, 6 females, same data asholotype except 6 June–20 July 1960 (AMNH).

Other material examined: USA: Nevada: Clark County: 1 K,Mercury, Nevada Test Site (36.94uN, 116.32uW), 6 June 1960,Atomic Energy Commission and Brigham Young University(AMNH); 1 K, same data except 11 June 1960 (AMNH); 1 L,same data except 20 June 1960 (AMNH); 1 K, same dataexcept 30 June 1960 (AMNH); 1 K, 22 June 1960 (AMNH); 1L, same data except 2 July 1960 (AMNH); 1 L, same dataexcept 8 July 1960 (AMNH); 1 L, same data except 11 July1960 (AMNH); 1 L, same data except 13 July 1960 (AMNH);L, same data except 25 July 1960 (AMNH).

Etymology.—The specific name refers to the explosion ofthe first nuclear device at the Nevada Test Site. It is to betreated as a noun in apposition.

Diagnosis.—This species is most similar to H. hanfordanaand H. banksi. It differs from H. hanfordana in the colorationof the chelicera with H. hanfordana dusky dark while H.kaboomi is dusky orange and from H. banksi in the shape ofthe cheliceral fixed finger. Hemerotrecha kaboomi has a ventralcurvature and a small hook apically while H. banksi has arounded tip. The ctenidia are longer than either of the abovespecies.

Description.—Male: chelicera dusky orange, dusky to darkmottled propeltidium (Fig. 21), and slightly lighter colored

54 THE JOURNAL OF ARACHNOLOGY

abdomen, leg I dusky on metatarsus only; legs II, III, IV duskyventrally.

Chelicera: fixed finger with a slight hook; the ventral apicalconcavity of the fixed finger shallow and extended, anteriortooth of movable finger triangular; posterior intermediatetooth separate from primary tooth; fondal notch obscure orabsent (Fig. 22).

Abdomen: ctenidia extending entire length of succeedingsternite (Fig. 24).

Palp: with 2–3 spine-like setae ectal ventrally (Fig. 23).

Male holotype: total length 10.5, cheliceral length 2.65,cheliceral width 1.12, propeltidium length 1.52, propeltidiumwidth 1.9, palpus length 7.0, first leg length 6.0, fourth leglength 12.5. Ratios: A/CP 6.12, chelicera width/fixed fingerwidth 4.48. Male paratypes (6): total length 8.5–10.0,

cheliceral length 2.16–2.40, cheliceral width 0.87–1.0, propel-tidium length 1.07–1.33, propeltidium width 2.02–2.40, palpuslength 4.0–6.5, first leg length 5.0–6.0, fourth leg length 8.5–9.5. Ratios: A/CP 5.18–5.97, cheliceral width/fixed fingerwidth 3.25–3.75.

Females: coloration as in males.

Chelicera: typical for the females of this group (Figs. 7, 13,24).

Genital region: typical of the group but with narrower armsand a more rounded posterior margin (Fig. 26).

Dimensions: Female allotype: total length 9.5, chelicerallength 3.0, cheliceral width 1.67, propeltidium length 1.18,propeltidium width 1.95, palpus length 5.0, first leg length 5.5,fourth leg length 6.5. Ratios: A/CP 4.07, genital operculumlength/genital operculum width 1.67.

Figures 15–20.—Hemerotrecha hanfordana, new species. 15–19. Male holotype: 15. Dorsal view, propeltidium; 16. Ectal view, right chelicerashowing anterior concavity (arrow); 17. Mesal view, right chelicera showing flagellum complex (arrow); 18. Mesal ventral view, right palpus; 19.Ventral view, ctenidia. 20. Female holotype, ectal view, right chelicera. Scale bar 5 1 mm.

BROOKHART & CUSHING—HEMEROTRECHA BANKSI GROUP (SOLIFUGAE) 55

Female paratypes (6): total length 9.5–11.5, chelicerallength 2.7–3.2, cheliceral width 0.9–1.67, propeltidium length1.18–1.45, propeltidium width 1.75–2.25, palpus length 5.0–5.5, first leg length 5.0–6.0, fourth leg length 6.5–7.5. Ratios:A/CP 4.0–4.4, genital operculum length/genital operculumwidth 1.4–2.0.

Remarks.—Females of H. kaboomi were collected early inJune, males in July, all in dry pitfalls as part of the survey ofthe Nevada Test Site. Allred et al. (1963) considered this areaas the boundary between the Mohave Desert and the GreatBasin geographic provinces.

Hemerotrecha marginata (Kraepelin 1911)

Figs. 27–30, 54

Eremobates marginatus Kraepelin 1911:103–105, figs. 4a–b.

Eremognatha marginata (Kraepelin): Roewer 1934:569, figs.116c, 128, 324p, 327c.

Hemerotrecha marginata (Kraepelin) Muma 1951:102, figs.198–201; Muma 1970:38, figs. 32–35.

Material examined.—Types: USA: California: Los AngelesCounty: male holotype, San Pedro (33.74uN, 118.29uW), 5June 1867 (ZSM, Roewer type #8376). Female allotype: same

data as holotype (ZSM). Paratypes: 1 male, 1 female, samedata as holotype (ZSM).

Other material.—USA: California: Inyo County: 1 L,Paxton Sand Dunes, 13 April–29 September 1982, E. Giuliani(ESS); Riverside County: 1 K, Banning (33.96uN, 116.89uW),April 1995, J.W. McSwain (ESS); San Bernardino County: 1L, Granite Cove (34.78uN, 115.66uW), Interstate 40 & KalbakeRoad, E. Fessler (UCR).

Diagnosis.—Hemerotrecha marginata is separated from H.californica, which it most closely resembles, by the colorationof chelicera, propeltidium, and eye tubercle, which are paleyellow. Palpus with metatarsus and distal 20% of tarsus dusky;ctenidia extend half way across succeeding sternite.

Description.—Male: Color: chelicera, propeltidium includ-ing eye tubercle, pale yellow. Palpus pale yellow withmetatarsus and distal 60% of tarsus dusky (Fig. 27); legsmostly pale, slightly dusky on ventral side of femur of legs III,IV, abdomen mottled, ctenidia extend halfway across suc-ceeding sternite.

Chelicera: typical for the group with only a slight apicalhook. Fixed finger regularly curved without teeth, shallow cuplike mesal ventral groove apically, movable finger with normal

Figures 21–26.—Hemerotrecha kaboomi, new species. 21–24. Male holotype: 21. Dorsal view, propeltidium; 22. Ectal view, chelicera; 23.Mesal ventral view, palpus; 24. Ventral view, ctenidia. 25–26. Female allotype: 25. Ectal view, right chelicera; 26. Ventral view, genitaloperculum. Scale bar 5 1 mm.

56 THE JOURNAL OF ARACHNOLOGY

primary tooth, flat pointed anterior tooth, 1–2 intermediateteeth separated from primary tooth. Base of fixed finger aswide as the base of movable finger. Fond indistinct, fondalteeth graded I, III, II, IV ectally and mesally (Fig. 28).

Abdomen: two thin ctenidia extending halfway acrosssucceeding sternite.

Palp: without spine-like setae (Fig. 27).

Dimensions: Male holotype: total length 10.0, chelicerallength 2.34, cheliceral width 1.21, propeltidium length 1.3,propeltidium width 1.7, palpus length 5.57, first leg length 5.5,fourth leg length 8.5. Ratios: A/CP 5.38, cheliceral width/fixedfinger width 3.46.

Males (4): total length 8.0–11.0, cheliceral length 2.1–2.45,cheliceral width 1.05–1.3, propeltidium length 0.8–1.5, pro-peltidium width 1.3–1.8, palpus length 4.0–6.2, first leg length4.5–6.1, fourth leg length 6.5–9.3. Ratios: A/CP 5.20–5.45,genital operculum length/genital operculum width 3.0–3.5.

Females: Coloration: same as in males.

Chelicera: typical for females of the group (Fig. 29).

Genital region: club shaped (Fig. 30).

Dimensions: Female allotype: total length 9.0, chelicerallength 2.3, cheliceral width 0.9, propeltidium length 1.2,propeltidium width 1.85, palpus length 5.0, first leg length 4.5,fourth leg length 7.0. Ratios: A/CP 4.70, genital operculumlength/genital operculum width 1.70.

Females (2): length 8.5–9.0, cheliceral length 2.2–2.7,cheliceral width 0.86–1.0, propeltidium length 1.05–1.46.propeltidium width 1.73–1.9, palpus length 4.0–5.4, first leglength 3.7–4.5, fourth leg length 5.8–7.4. Ratios: A/CP 3.6–4.7,genital operculum length/genital operculum width 1.67–1.70.

Remarks.—The illustration of the ctenidia provided byMuma (1951) was based on the drawing published byKraepelin (1911). Muma (1970) re-drew the ctenidia basedupon examination of the type specimen. In both the 1951 andthe 1970 drawings, Muma illustrates three ctenidia found onthe type specimen. However, in the 1970 publication, Mumaassumed that it possessed four ctenidia, and that one hadsimply broken off. The males that we examined consistentlyhad but two ctenidia as do all others of this group.

Hemerotrecha truncata Muma 1951

Figs. 31–36, 54

Hemerotrecha truncata Muma 1951:102, fig. 197; Muma1970:41.

Material examined.—Type: USA: California: Tulare Coun-ty: male holotype, Exeter (36.3uN, 119.14uW), 16 May 1909,C. L. Fox (AMNH).

Other material: USA: California: Inyo County: 1 L,Whippoorwill Canyon (37.00uN, 117.94uW), May 1983–June1984, D.Giuliani (CA); Stanislaus County: 2 L, 1 K, Lagrange

Figures 27–30.—Hemerotrecha marginata (Kraepelin). 27–28. Male holotype: 27. Mesal ventral view, right palpus; 28. Ectal view, leftchelicera. 29–30. Female allotype: 29. Ectal view, right chelicera; 30. Ventral view, genital operculum. Scale bars 5 1 mm.

BROOKHART & CUSHING—HEMEROTRECHA BANKSI GROUP (SOLIFUGAE) 57

(37.67uN, 120.46uW), 27 May 1976, J. Collins (CAS); 2 L, 1 K,Turlock (37.5uN, 120.85uW), 2 June 1976, J. Collins (CAS);Tulare County: 1 L, 1 K, Kaweah (36.47uN, 118.92uW), 13 May1963, J. Boswell (ESS).

Diagnosis.—This species is easily distinguished by theparrot-beak shaped hook on the tip of male fixed finger.Specimens are darkly colored.

Description.—Males: Color: chelicera dusky to dark yellow,propeltidium and abdomen chocolate brown (Fig. 31), palpaltarsus, metatarsus, tip of tibia darkly colored (Fig. 32); all legsdark to dusky brown; eye tubercle dark.

Chelicera: fixed finger sharply hooked (parrot beak) withdeep cup-like mesal ventral groove apically, no dorsal trough/

ridge. Dentition typical of the group with posterior interme-diate tooth of movable finger separate from primary tooth(Fig. 32).

Abdomen: two long, thin ctenidia extending the length ofthe succeeding sternite (Fig 34).

Palp: as in Fig. 33.

Dimensions: Male holotype: total length 11.0, chelicerallength 3.1, cheliceral width 1.4, propeltidium length 2.01,propeltidium width 2.8, palpus length 9.0, first leg length 7.0,fourth leg length 13.5. Ratios: A/CP 5.77, cheliceral width/fixed finger 4.24.

Male paratypes (7): total length 7.5–11.0, cheliceral length2.0–2.5, cheliceral width 0.8–1.5, propeltidium length 0.9–2.0,

Figures 31–36.—Hemerotrecha truncata Muma. 31–34. Male holotype: 31. Dorsal view, propeltidium; 32. Ectal view, right chelicera; 33.Mesal ventral view, right palpus; 34. Ventral view, ctenidia. 35–36. Female: 35. Ectal view, right chelicera; 36. Ventral view, genital operculum.Scale bar 5 1 mm.

58 THE JOURNAL OF ARACHNOLOGY

propeltidium width 1.5–2.8, palpus length 6.5–9.0, first leglength 4.5–7.0, fourth leg length 6.5–9.0. Ratios: A/CP 5.7–6.8,cheliceral width/fixed finger width 2.8–4.2.

Female: coloration as in the male.

Chelicera: typical of the group except posterior intermediatetooth of the movable finger is on the notch of the primarytooth (Fig. 35);

Palp: three pair of spine-like setae on ventral margin ofmetatarsus.

Genital region: typical of the group (Fig. 36).

Female measurements (2): total length 12.0–13.0, chelicerallength 3.3–3.6, cheliceral width 1.35–1.46, propeltidium length1.44–1.86, propeltidium width 2.5–2.6, palpus length 7.0–8.0,first leg length 6.5–7.0, fourth leg length 12.5–14.5. Ratios: A/CP 5.0–5.7, genital operculum length/genital operculum width1.7–1.8.

Remarks.—This very distinctive member of the H. banksigroup inhabits the Mohave Desert Region of easternCalifornia.

Hemerotrecha pseudotruncata new species

Figs. 37–42, 54

Material examined.—USA: Nevada: Nye County: maleholotype Monitor Summit (38.8uN, 115.5uW), 27.3 km E. of

Tonapah, October 1982–September 1983, D. Giuliani (ESS);female allotype: Currant Summit, 16.1 km E. of Currant,October 1982–September 1983, D. Giuliani (ESS).

Other material: USA: California: Inyo County: China Lake3 L, 1 K, Death Valley Deep Springs (37.37uN, 117.98uW), May1983–June 1984, D. Giuliani (CAS); 1 L, Fish Lake Valley,Waucoba Spring (36.7uN, 116.84uW), May–October 1983, D.Giuliani (CAS); 3 L, 1 K, Saline Range (coordinates notpossible), March 1979–May 1983, D. Giuliani (CAS); Nevada:Esmeralda County 1 L, 8 km SW Lida (37.321uN, 117.8uW);Nye County: 1 L, 1 K, Grapevine Mountains, Phinney Canyon(36.99uN, 117.03uW), March–September 1983, D. Giuliani(CAS); 1 L, Monitor Summit (38.05uN, 117.22uW), October1982–September 1983, D. Giuliani (CAS); 1 L, 1 K, CurrantSummit (38.74uN, 115.47uW), March–September 1983, D.Giuliani (CAS).

Etymology.—The species name refers to the modified hookat the tip of male fixed finger, which is similar to H. truncatabut not as radically modified.

Diagnosis.—This species differs from H. truncata by havingless radical hook-like tip on the male fixed finger, slightlybroader and shorter ctenidia, and slightly lighter coloration.

Description.—Male: chelicera dusky orange, propeltidiumdusky (Fig. 37), palpi usually dark to dusky on apical 80% of

Figures 37–42.—Hemerotrecha pseudotruncata, new species. 37–40. Male holotype: 37. Dorsal view, propeltidium; 38. Ectal view, leftchelicera; 39. Dorsal view, right palpus; 40. Ventral view, ctenidia. 41–42. Female allotype: 41. Ectal view, right chelicera; 42. Dorsal view, genitaloperculum. Scale bar 5 1 mm.

BROOKHART & CUSHING—HEMEROTRECHA BANKSI GROUP (SOLIFUGAE) 59

metatarsus (Fig. 39), eye tubercle dark, legs dark to duskyventrally.

Chelicera: with sharp hook apically but not nearly asprominent as in H. truncata, a short, deep concave cup-likemesal ventral groove apically, anterior ventral edge concave,movable finger with large primary tooth, smaller anteriortooth, single intermediate tooth separated from primary tooth,crenulations on the anterior dorsal edge; obscure to smallfondal notch (Fig. 38).

Abdomen: two pointed ctenidia extending about one halfthe length of succeeding sternite (Fig. 40).

Palp: 2–3 spine-like setae on ventral metatarsus (Fig. 39).

Male holotype: total length 8.5, cheliceral length 2.1,cheliceral width 1.0, propeltidium length 1.1, propeltidiumwidth 1.4, palpus length 8.6, first leg length 5.5, fourth leglength 7.0. Ratios: A/CP 6.6, cheliceral width/fixed fingerwidth 3.78.

Male paratypes (n58): total length 8.0–10.0, chelicerallength 1.9–2.7, cheliceral width 0.76–1.0, propeltidiumlength 1.0–1.36, propeltidium width 1.3–1.8, palpus length8.0–10.5, first leg length 5.0–6.5, fourth leg length 6.0–7.0.Ratios: A/CP 6.4–6.5, cheliceral width/fixed finger width 3.33–4.1.

Female: coloration as in males.

Chelicera: typical for females of this group (Fig. 41).

Genital region: smaller and more oval than other membersof this group (Fig. 42).

Dimensions: Female allotype: total length 8.0, chelicerallength 2.6, cheliceral width 0.9, propeltidium length 1.33,propeltidium width 1.76, palpus length 5.0, first leg length 5.0,fourth leg length 6.5. Ratios: A/CP 3.1, genital operculumlength/genital operculum width 0.8.

Female (n54): total length 8.0–8.5, cheliceral length 2.8–3.0,cheliceral width 1.0, propeltidium length 1.4–1.55, propelti-dium width 1.65–1.8, palpus length 5.5–5.8, first leg length5.5–6.0, fourth leg length 6.5–7.0. A/CP 3.8–4.0, genitaloperculum length/genital operculum width 0.8–0.9.

Remarks.—Hemerotrecha pseudotruncata seems closely re-lated to H. truncata.

Hemerotrecha vetteri new species

Figs. 43–47, 54

Material examined.—Type: USA: California: Santa BarbaraCounty: male holotype, Vandenburg Air Force Base (34.76uN,120.46uW), 30 April 2004, Abela, Pierce, Pratt (DMNS).

Etymology.—This species is named for the inveterateinvertebrate collector and arachnologist, Rick Vetter, whosent this and many other specimens for examination.

Figures 43–47.—Hemerotrecha vetteri, new species, male holotype: 43. Dorsal view, propeltidium; 44. Ectal view, right chelicera (arrowdenotes dorsal ridge); 45. Mesal view, right chelicera (arrow denotes flagella complex); 46. Mesal ventral view, right palpus; 47. Ventral view,ctenidia. Scale bar 5 1 mm.

60 THE JOURNAL OF ARACHNOLOGY

Diagnosis.—This totally dark species is most similar to H.banksi except for its completely dark coloration. The presenceof the dorsal, trough-like structure on the cheliceral fixedfinger is unique to the group. Females unknown.

Description.—Male: Color: total body dark with no lightpatches (Fig. 43). Appendages dark over entire length.

Chelicera: similar to H. banksi with a short, small ventralconcavity anteriorly. A highly visible ridge on the distal,ventral margin of fixed finger. Fondal notch absent orobscure, posterior intermediate tooth of movable fingerseparate from primary tooth (Figs. 44, 45).

Abdomen: ctenidia short, pointed, and extending abouthalfway across succeeding abdominal segment (Fig. 47).

Palp: several spine-like setae on mesal ventral side (Fig. 46).Male holotype: total length 11.0, cheliceral length 2.7,

cheliceral width 1.2, propeltidium length 1.4, propeltidiumwidth 2.1, palpus length 7.0, first leg length 4.5, fourth leg

length 10.0. Ratios: A/CP 6.14, cheliceral width/fixed fingerwidth 3.2.

Remarks.—No females are known at present.

Hemerotrecha prenticei new species

Figs. 48–52, 54

Material examined.—USA: California: male holotype fromSouthwest Riverside County Multispecies Reserve (LakeSkinner), Riverside County (33.38uN, 117.00uW), 9–12 May2000, Tom Prentice (DMNS). Female allotype: collected fromsame site, 15–18 May 2000 (DMNS). Paratypes: 4 males, 2females, collected from same site, various dates (DMNS); 4males, 1 female, collected from same site, various dates(UCR).

Other material: USA: California: Kern County, 1L, 1K,Dove Springs (35.42uN, 118.00uW) June 2003, collected by theUnited States Geological Survey, San Diego Office; 4 L, same

Figures 48–53.—Hemerotrecha prenticei, new species. 48–51. Male holotype: 48. Dorsal view, propeltidium; 49. Dorsal view, abdomen; 50.Dorsal view, right palpus; 51. Ectal view, right chelicera. 52. Ventral view, male ctenidia. 53. Ectal view, female chelicera. Scale bars 5 1 mm.

BROOKHART & CUSHING—HEMEROTRECHA BANKSI GROUP (SOLIFUGAE) 61

data except May 2004 (DMNS); 10 L, 4 K, Elk Hills (35.16uN,119.31uW), T. Prentice (DMNS, UCR); 2 L, 3 K, El PasoMountains (35.26uN, 117.49uW) August 1965, D. Gibo(UCN); 1 L, El Paso Mountains (35.26uN, 117.49uW) 24 June1970, Wendel Icenogle (UCN).

Etymology.—Named for the collector, Tom Prentice,University of California, Riverside.

Diagnosis.—This species differs from the closely related H.pseudotruncata in its darker overall coloration and thebroader, shorter ctenidia.

Description.—Male: Color: base coloration a creamy, darkorange, propeltidium with lateral dusky areas, eye tubercledark, abdomen darker dorsally and ventrally with a medianwhite stripe dorsally extending from propeltidium to sixthabdominal segment (Figs. 48, 49), palpal metatarsus dark overthe entire length (Fig. 50), all legs dusky to dark on tarsus,metatarsus, tibia, and femur, coxa lighter. Leg I lighterventrally. Malleoli edges dark.

Chelicera: fixed finger with rounded tip and typical cup likemesal ventral groove apically. No recurved area apically.Movable finger with primary tooth, two intermediate teeth,and an anterior tooth, Posterior intermediate tooth separatefrom the primary tooth. Fondal teeth graded I, III, II, IV

ectally and I, III, II mesally. No apparent fondal notch(Fig. 51). Flagella typical of the group.

Abdomen: 2 short, broad, pointed ctenidia (Fig. 52). Theseare the shortest of any in this species group.

Palp: entire palp covered with setae but no spine-like setaepresent (Fig. 50).

Dimensions: Male holotype: total length 10.5, chelicerallength 2.2, cheliceral width 0.82, propeltidium length 1.35,propeltidium width 1.84, palpus length 6, first leg length 4.5,fourth leg length 8. Ratios: A/CP 5.25, cheliceral width/fixedfinger width 3.0.

Male paratypes (n55): total length 9.0–11.0, chelicerallength 2.0–2.4, cheliceral width 0.8–1.0, propeltidium length1.2–1.4, propeltidium width 1.6–1.9, palpus length 9.0–11.0,first leg length 4.5–5.5, fourth leg length 7.0–9.5. Ratios: A/CP5.0–5.9, cheliceral width/fixed finger width 3.0–4.3.

Female allotype: color, same as the male.

Chelicera: fixed finger typical of the group but on themovable finger the posterior intermediate tooth is closelyoppressed to the posterior tooth which differs from otherfemales of this group (Fig. 53).

Palp: same as the male with 1–2 spine-like setae ventrally.

Genital operculum: typical for the group.

Figure 54.—Distribution map of Hemerotrecha banksi species group in the Western USA.

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Dimensions: Female allotype: total length 10.5, chelicerallength 2.25, cheliceral width 1.05, propeltidium length 1.2,propeltidium width 1.75, palpal length 7.0, first leg length 5.0,fourth leg length 7.5. Ratios: A/CP 5.27, genital length/genitalwidth 2.0.

Female paratypes (n55): total length 10.5–11.5, chelicerallength 2.3–3.1, cheliceral width 0.9–1.0, propeltidium length1.2–1.3, propeltidium width 1.7–2.2, palpus length 5.0–7.0,first leg length 4.0–5.0, fourth leg length 6.5–7.0. Ratios: A/CP4.3–5.7, genital operculum length/ genital operculum width2.3–3.1.

DISCUSSION

Adult members of the Hemerotrecha banksi group havebeen collected as early as the first week in May and as late asthe middle of October. A total of 82 males and 27 females wereexamined in this study, indicating a greater vagility among themales. Several vials contained information suggesting that thecollected specimens were active during the daytime. Wesuggest that all of the species of this group are diurnal. Forsuch diurnal species, color ornamentation, particularly of thepalps, could be important during courtship and mating.Although mating patterns of solifuges remain largely un-known (Punzo 1998), the few descriptions available (Muma1966; Wharton 1981; Punzo 1998) involve the presentation ofpalps by both sexes.

Most North American Solifugae have white to pale malleoli.The presence of the dark edged malleoli in the H. banski groupwas not noted by Muma (1951, 1970) although he did identifythem in other species (Muma 1951, 1962); i.e., Eremochelisrothi (Muma 1962), Eremochelis larreae (Muma 1962),Hemerotrecha bidepressus (Muma 1951), and (Eremochelisarcellus Muma 1962). Examination of the types of thesespecies showed at most a very tiny edge of black. He also didnot describe the cup-like cavity on the tip of the male fixedfinger as a mesal ventral groove, which is surprising since hedid so in the Eremochelis branchi group (Muma 1970).Eremochelis striodorsalis (Muma 1962) has an obvious ridgewhich Muma (1962) called a mesal dorsal groove. Examina-tion of the type of E. striodorsalis reveals a faint mesal ventralgroove. The dorsal ventral ridge of E. striodorsalis and H.vetteri are very similar.

ACKNOWLEDGMENTS

We would like to thank all of the curators, collectionmanagers, and collectors who kindly provided us withmaterial for this study. Individuals include Lorenzo Prendini(AMNH); Richard Bauman (BYU); Charles Griswold (CAS);Robert Hogue (CSN); Tom Prentice (UCR), Cheryl Barr(ESS); G.B. Edwards (FSCA); Gonzalo Giribet and LauraLeibensperger (MCZ); Paisley Cato and Jim Berrian (SMNH);R. Zack (WSU); and Roland Melzer (ZSM). Thanks also toWendell Icenogle, Winchester, California for loaning speci-mens used in this study. Specimens from the Denver Museumof Nature & Science (DMNS) were also used.

Thanks also to Joey Slowik and Aaron Spriggs forassistance with photography and in compiling distributionmaps and illustrations. This work was supported by NationalScience Foundation grants DBI-0346378 and DBI-0640245awarded to P.E.C.

LITERATURE CITED

Allred, D.M., D.E. Beck & C.D. Jorgensen. 1963. Biotic communitiesof the Nevada Test Site. Brigham Young University ScienceBulletin, Biological Series 2(2):1–52.

Banks, N. 1899. A new solpugid from California. Proceedings of theEntomological Society of Washington 4:314–315.

Banks, N. 1900. Synopses of North American invertebrates. IX. Thescorpions, solpugids, and pedipalpi. American Naturalist34:421–427.

Banks, N. 1903. A new genus of Solpugida. Entomological News14:78–79.

Banks, N. 1904. Some Arachnida from California. Proceedings of theCalifornia Academy of Science 3:331–374.

Brookhart, J.O. & I.P. Brookhart. 2006. An annotated checklist ofNorth American Solifugae with type depositories, abundance, andnotes on their zoogeography. Journal of Arachnology 34:299–329.

Brookhart, J.O. & P.E. Cushing. 2002. New species of Eremobatidae(Arachnida, Solifugae) from North America. Journal of Arachnol-ogy 30:84–97.

Brookhart, J.O. & P.E. Cushing. 2004. The systematics of theEremobates scaber group (Solifugae, Eremobatidae). Journal ofArachnology 32:284–312.

Brookhart, J.O. & M.H. Muma. 1981. The pallipes species-group ofEremobates Banks (Solpugida: Arachnida). Florida Entomologist64:283–308.

Brookhart, J.O. & M.H. Muma. 1987. Arenotherus, a new genus ofEremobatidae (Solpugida) in the United States, pp. 1–18 and 4plates. Printed for the authors by Cherry Creek High School PrintShop, Englewood, Colorado.

Cloudsley-Thompson, J.L. 1958. Spiders, Scorpions, Centipedes andMites. Pergamon Press, Oxford, UK. 278 pp.

Harvey, M.S. 2003. Catalogue of the Smaller Arachnid Orders of theWorld: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinuleiand Solifugae. CSIRO Publishing, Collingwood, Victoria, Aus-tralia. 385 pp.

Kraepelin, K. 1911. Beitrage zur Systematik der Gliederspinnen.Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten /Mitteilungen aus dem Naturhistorischen Museum in Hamburg28(2):57–107.

Lawrence, R.F. 1960. Some observations on the arachnid fauna ofSouth West Africa. South African Museums Association Bulletin7:67–72.

Lawrence, R.F. 1962. Solifuges, scorpions and Chilopoda of theNamib Desert. Annals of the Transvaal Museum 24:213–222.

Maury, E.A. 1985. Las familias de solifugos Americanos y sudistribucion geographica (Arachnida, Solifugae). Physis, BuenosAires 42:73–80.

Muma, M.H. 1951. The arachnid order Solpugida in the UnitedStates. Bulletin of the American Museum of Natural History97:35–141.

Muma, M.H. 1962. The arachnid order Solpugida in the UnitedStates, Supplement 1. American Museum Novitates 2092:1–44.

Muma, M.H. 1963. Solpugida of the Nevada Test Site. BYU ScienceBulletin. Biology. Series 3(2):1–15.

Muma, M.H. 1966. Mating behavior in the solpugid genusEremobates Banks. Animal Behaviour 14:346–350.

Muma, M.H. 1970. A synoptic review of North American, CentralAmerican and West Indian Solpugida (Arthropoda: Arachnida).Arthropods of Florida and Neighboring Land Areas 5:1–62.

Muma, M.H. 1989. New species and records of Solpugida (Arach-nida) from the United States. Published for the author by DouglasPrint Shop, Douglas, Arizona. 50 pp., 8 plates.

Muma, M.H. & J.O. Brookhart. 1988. The Eremobates palpisetulosusspecies-group (Solpugida: Eremobatidae) in the United States, pp.1–65, plates 1–20. Published for the authors by Cherry Creek HighSchool Print Shop, Englewood, Colorado.

BROOKHART & CUSHING—HEMEROTRECHA BANKSI GROUP (SOLIFUGAE) 63

Punzo, F. 1998. The Biology of Camel Spiders (Arachnida,Solifugae). Kluwer Academic Publishers, Norwell, Massachusetts.301 pp.

Roewer, C.F.R. 1934. Solifugae, Palpigradi. Pp. 481–608. In Klassenund Ordnungen des Tierreichs. (H.G. Bronn, ed.). Volume 5:Arthropoda, Division 4: Arachnoidea, Book 4. AkademischeVerlagsgesellschaft M.B.H., Leipzig.

Wharton, R.A. 1981. Namibian Solifugae (Arachnida). CimbesiaMemoir 5:1–87.

Wharton, R.A. 1987. Biology of the diurnal Metasolpuga picta(Kraepelin) (Solifugae, Solpugidae) compared with that ofnocturnal species. Journal of Arachnology 14:363–383.

Xavier, R. & L.S. Rocha. 2001. Autoecology and description ofMummucia mauryi (Solifugae, Mummuciidae) a new solifugefrom Brazilian semi-arid caatinga. Journal of Arachnology29:127–134.

Manuscript received 19 February 2007, revised 7 December 2007.

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